Trade-Offs in the Sensory Brain between Diurnal and Nocturnal Rodents.

Introduction: Transitions in temporal niche have occurred many times over the course of mammalian evolution. These are associated with changes in sensory stimuli available to animals, particularly with visual cues, because levels of light are so much higher during the day than at night. This relationship between temporal niche and available sensory stimuli elicits the expectation that evolutionary transitions between diurnal and nocturnal lifestyles will be accompanied by modifications of sensory systems that optimize the ability of animals to receive, process, and react to important stimuli in the environment.

Measuring salivary cortisol in wild carnivores.

Salivary hormone analyses provide a useful alternative to fecal and urinary hormone analyses in non-invasive studies of behavioral endocrinology. Here, we use saliva to assess cortisol levels in a wild population of spotted hyenas (Crocuta crocuta), a gregarious carnivore living in complex social groups. We first describe a novel, non-invasive method of collecting saliva from juvenile hyenas and validate a salivary cortisol assay for use in this species.

Sex Differences in Spotted Hyenas.

The apparent virilization of the female spotted hyena raises questions about sex differences in behavior and morphology. We review these sex differences to find a mosaic of dimorphic traits, some of which conform to mammalian norms. These include space-use, dispersal behavior, sexual behavior, and parental behavior.

Early-life social experience affects offspring DNA methylation and later life stress phenotype.

Studies in rodents and captive primates suggest that the early-life social environment affects future phenotype, potentially through alterations to DNA methylation. Little is known of these associations in wild animals. In a wild population of spotted hyenas, we test the hypothesis that maternal care during the first year of life and social connectedness during two periods of early development leads to differences in DNA methylation and fecal glucocorticoid metabolites (fGCMs) later in life.

Circadian and photic modulation of daily rhythms in diurnal mammals.

The temporal niche that an animal occupies includes a coordinated suite of behavioral and physiological processes that set diurnal and nocturnal animals apart. The daily rhythms of the two chronotypes are regulated by both the circadian system and direct responses to light, a process called masking. Here we review the literature on circadian regulations and masking responses in diurnal mammals, focusing on our work using the diurnal Nile grass rat (Arvicanthis niloticus) and comparing our findings with those derived from other diurnal and nocturnal models.

Distributions of GABAergic and glutamatergic neurons in the brains of a diurnal and nocturnal rodent.

Light influences the daily patterning of activity by both synchronizing internal clocks to environmental light-dark cycles and acutely modulating arousal states, a process known as masking. Masking responses are completely reversed in diurnal and nocturnal species. In nocturnal rodents, masking is mediated through a subset of intrinsically photosensitive retinal ganglion cells (ipRGCs) whose projections are similar in diurnal and nocturnal rodents.

The Cost of Activity during the Rest Phase: Animal Models and Theoretical Perspectives.

For humans, activity during the night is correlated with multiple pathologies that may reflect a lack of harmony among components of the circadian system; however, it remains difficult to identify causal links between nocturnal activity and different pathologies based on the data available from epidemiological studies. Animal models that use forced activity or timed sleep deprivation provide evidence of circadian disruptions that may be at the core of the health risks faced by human night and shift workers. One valuable insight from that work is the importance of changes in the distribution of food intake as a cause of metabolic imbalances associated with activity during the natural rest phase.

Normal behavioral responses to light and darkness and the pupillary light reflex are dependent upon the olivary pretectal nucleus in the diurnal Nile grass rat.

The olivary pretectal nucleus (OPT) is a midbrain structure that receives reciprocal bilateral retinal projections, is involved in the pupillary light reflex, and connects reciprocally with the intergeniculate leaflet (IGL), a retinorecipient brain region that mediates behavioral responses to light pulses (i.e., masking) in diurnal Nile grass rats.

Suprachiasmatic Nucleus and Subparaventricular Zone Lesions Disrupt Circadian Rhythmicity but Not Light-Induced Masking Behavior in Nile Grass Rats.

The ventral subparaventricular zone (vSPVZ) receives direct retinal input and influences the daily patterning of activity in rodents, making it a likely candidate for the mediation of acute behavioral responses to light (i.e., masking).

Central melanopsin projections in the diurnal rodent, Arvicanthis niloticus.

The direct effects of photic stimuli on behavior are very different in diurnal and nocturnal species, as light stimulates an increase in activity in the former and a decrease in the latter. Studies of nocturnal mice have implicated a select population of retinal ganglion cells that are intrinsically photosensitive (ipRGCs) in mediation of these acute responses to light. ipRGCs are photosensitive due to the expression of the photopigment melanopsin; these cells use glutamate and pituitary adenylate cyclase-activating polypeptide (PACAP) as neurotransmitters.

Acute effects of light on the brain and behavior of diurnal Arvicanthis niloticus and nocturnal Mus musculus.

Photic cues influence daily patterns of activity via two complementary mechanisms: (1) entraining the internal circadian clock and (2) directly increasing or decreasing activity, a phenomenon referred to as "masking". The direction of this masking response is dependent on the temporal niche an organism occupies, as nocturnal animals often decrease activity when exposed to light, while the opposite response is more likely to be seen in diurnal animals. Little is known about the neural mechanisms underlying these differences.

Intergeniculate leaflet lesions result in differential activation of brain regions following the presentation of photic stimuli in Nile grass rats.

The intergeniculate leaflet (IGL) plays an important role in the entrainment of circadian rhythms and the mediation of acute behavioral responses to light (i.e., masking).

Behavioral Masking and cFos Responses to Light in Day- and Night-Active Grass Rats.

Light not only entrains the circadian system but also has acute effects on physiology and behavior, a phenomenon known as masking. Behavioral masking responses to bright light differ in diurnal and nocturnal species, such that light increases arousal in the former and decreases it in the latter. Comparisons made within a species that displays both diurnal and nocturnal patterns of behavior may provide insight into how masking differs between chronotypes and the association between mechanisms controlling masking and the circadian drive for activity.

Lesions of the Intergeniculate Leaflet Lead to a Reorganization in Circadian Regulation and a Reversal in Masking Responses to Photic Stimuli in the Nile Grass Rat.

Light influences the daily patterning of behavior by entraining circadian rhythms and through its acute effects on activity levels (masking). Mechanisms of entrainment are quite similar across species, but masking can be very different. Specifically, in diurnal species, light generally increases locomotor activity (positive masking), and in nocturnal ones, it generally suppresses it (negative masking).

Pregnancy affects FOS rhythms in brain regions regulating sleep/wake state and body temperature in rats.

Circadian rhythms in behavior and physiology change substantially as female mammals undergo the transition from a non-pregnant to a pregnant state. Here, we examined the possibility that site-specific changes in brain regions known to regulate the sleep/wake cycle and body temperature might reflect altered rhythms in these overt functions. Specifically, we compared daily patterns of immunoreactive FOS in early pregnant and diestrous rats in the medial septum (MS), vertical and horizontal diagonal bands of Broca (VDB and HDB), perifornical lateral hypothalamus (LH), and ventrolateral, medial, and median preoptic areas (VLPO, MPA, and MnPO, respectively).

Dim nighttime light impairs cognition and provokes depressive-like responses in a diurnal rodent.

Circadian disruption is a common by-product of modern life. Although jet lag and shift work are well-documented challenges to circadian organization, many more subtle environmental changes cause circadian disruption. For example, frequent fluctuations in the timing of the sleep/wake schedule, as well as exposure to nighttime lighting, likely affect the circadian system.

Acute behavioral responses to light and darkness in nocturnal Mus musculus and diurnal Arvicanthis niloticus.

The term masking refers to immediate responses to stimuli that override the influence of the circadian timekeeping system on behavior and physiology. Masking by light and darkness plays an important role in shaping an organism's daily pattern of activity. Nocturnal animals generally become more active in response to darkness (positive masking) and less active in response to light (negative masking), and diurnal animals generally have opposite patterns of response.

Site-specific changes in brain extra-SCN oscillators during early pregnancy in the rat.

The influence of circadian timekeeping systems on behavior and physiology can change substantially as female mammals undergo the transition from a nonpregnant to a pregnant state. Here, we examined the possibility that site-specific changes in extra-SCN oscillators and in local rhythms might coincide with the emergence of new patterns of temporal organization among various behavioral and physiological rhythms. Specifically, we compared daily patterns of immunoreactive FOS and PER2 in 3 brain regions of pregnant and diestrous rats.

Projections of the suprachiasmatic nucleus and ventral subparaventricular zone in the Nile grass rat (Arvicanthis niloticus).

The phases of many circadian rhythms differ between diurnal and nocturnal species. However, rhythms within the hypothalamic suprachiasmatic nucleus (SCN), which contains the central circadian pacemaker, are very similar, suggesting that the mechanisms underlying phase preference lie downstream of the SCN. Rhythms in Fos expression in the ventral subparaventricular zone (vSPVZ), a major target of the SCN, differ substantially between diurnal Nile grass rats and nocturnal lab rats, raising the possibility that the vSPVZ modulates the effects of SCN signals at its targets.

PER2 rhythms in the amygdala and bed nucleus of the stria terminalis of the diurnal grass rat (Arvicanthis niloticus).

The suprachiasmatic nucleus (SCN) of the hypothalamus is the central pacemaker that controls circadian rhythms in mammals. In diurnal grass rats (Arvicanthis niloticus), many functional aspects of the SCN are similar to those of nocturnal rodents, making it likely that the difference in the circadian system of diurnal and nocturnal animals lies downstream from the SCN. Rhythms in clock genes expression occur in several brain regions outside the SCN that may function as extra-SCN oscillators.

The substructure of the suprachiasmatic nucleus: Similarities between nocturnal and diurnal spiny mice.

Evolutionary transitions between nocturnal and diurnal patterns of adaptation to the day-night cycle must have involved fundamental changes in the neural mechanisms that coordinate the daily patterning of activity, but little is known about how these mechanisms differ. One reason is that information on these systems in very closely related diurnal and nocturnal species is lacking. In this study, we characterize the suprachiasmatic nucleus (SCN), the primary brain structure involved in the generation and coordination of circadian rhythms, in two members of the genus Acomys with very different activity patterns, Acomys russatus (the golden spiny mouse, diurnal) and Acomys cahirinus (the common spiny mouse, nocturnal).

Masking and temporal niche switches in spiny mice.

Activity patterns are the product of interactions between an internal circadian clock and direct responses to photic and nonphotic features of the environment that are said to "mask" the influence of that clock. Evolutionary transitions between nocturnality and diurnality involve changes in mechanisms underlying both of these processes. Here, the authors examined how masking influences activity patterns of golden spiny mice ( Acomys russatus), which can be either nocturnal or diurnal, and common spiny mice (Acomys cahirinus), which are strictly nocturnal.

Changing patterns of daily rhythmicity across reproductive states in diurnal female Nile grass rats (Arvicanthis niloticus).

A suite of changes in circadian rhythms have been described in nocturnal rodents as females go through pregnancy and lactation, but there is no information on such patterns in diurnal species. As the challenges faced by these two groups of animals are somewhat different, we characterized changes in activity and core body temperature (T(b)) in female diurnal Nile grass rats (Arvicanthis niloticus) as they went through a series of reproductive states: virgin, pregnant, pregnant and lactating, lactating only, and post-weaning. The phase of neither rhythm varied, but the amplitude did.

Daily immediate early gene expression in the suprachiasmatic nucleus of male and female Octodon degus.

Diurnal and nocturnal species have different patterns of general activity, sleep-wake rhythms, and endocrine rhythms, but the mechanisms underlying these differences are not clear. Here, we tested the hypothesis that rhythms in immediate early gene (IEG) products in the suprachiasmatic nucleus (SCN) of a diurnal rodent (Octodon degus) reflect their diurnal chronotype. We also compared male and female degus with respect to temporal patterns of expression of one of these gene products, Fos-related Antigen (FRA).

Plasticity of circadian activity and body temperature rhythms in golden spiny mice.

Most animals can be categorized as nocturnal, diurnal, or crepuscular. However, rhythms can be quite plastic in some species and vary from one individual to another within a species. In the golden spiny mouse (Acomys russatus), a variety of rhythm patterns have been seen, and these patterns can change considerably as animals are transferred from the field into the laboratory.